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<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Hum. Neurosci.</journal-id>
<journal-title>Frontiers in Human Neuroscience</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Hum. Neurosci.</abbrev-journal-title>
<issn pub-type="epub">1662-5161</issn>
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<publisher-name>Frontiers Media S.A.</publisher-name>
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<article-id pub-id-type="doi">10.3389/fnhum.2024.1401494</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Human Neuroscience</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Dyadic body competence predicts movement synchrony during the mirror game</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Moffat</surname> <given-names>Ryssa</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
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<contrib contrib-type="author">
<name><surname>Roos</surname> <given-names>Leonie</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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<contrib contrib-type="author">
<name><surname>Casale</surname> <given-names>Courtney</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
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<contrib contrib-type="author" corresp="yes">
<name><surname>Cross</surname> <given-names>Emily S.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
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<aff id="aff1"><sup>1</sup><institution>Professorship for Social Brain Sciences, ETH</institution>, <addr-line>Zurich</addr-line>, <country>Switzerland</country></aff>
<aff id="aff2"><sup>2</sup><institution>Institute of Cognitive Science, Osnabr&#x00FC;ck University</institution>, <addr-line>Osnabr&#x00FC;ck</addr-line>, <country>Germany</country></aff>
<aff id="aff3"><sup>3</sup><institution>School of Psychological Sciences, Macquarie University</institution>, <addr-line>Sydney, NSW</addr-line>, <country>Australia</country></aff>
<author-notes>
<fn fn-type="edited-by" id="fn0002">
<p>Edited by: Jason Friedman, Tel Aviv University, Israel</p>
</fn>
<fn fn-type="edited-by" id="fn0003">
<p>Reviewed by: Juha M. Lahnakoski, LVR Klinik D&#x00FC;sseldorf, Germany</p>
<p>Hila Zahava Gvirts, Ariel University, Israel</p>
</fn>
<corresp id="c001">&#x002A;Correspondence: Ryssa Moffat, <email>ryssa.moffat@gess.ethz.ch</email>; Emily S. Cross, <email>emily.cross@gess.ethz.ch</email></corresp>
</author-notes>
<pub-date pub-type="epub">
<day>19</day>
<month>06</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>18</volume>
<elocation-id>1401494</elocation-id>
<history>
<date date-type="received">
<day>15</day>
<month>03</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>07</day>
<month>06</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2024 Moffat, Roos, Casale and Cross.</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Moffat, Roos, Casale and Cross</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>The process of synchronizing our body movements with others is known to enhance rapport, affect, and prosociality. Furthermore, emerging evidence suggests that synchronizing activities may enhance cognitive performance. Unknown, by contrast, is the extent to which people&#x2019;s individual traits and experiences influence their ability to achieve and maintain movement synchrony with another person, which is key for unlocking the social and affective benefits of movement synchrony. Here, we take a dyad-centered approach to gain a deeper understanding of the role of embodiment in achieving and maintaining movement synchrony. Using existing data, we explored the relationship between body competence and body perception scores at the level of the dyad, and the dyad&#x2019;s movement synchrony and complexity while playing a 2.5-min movement mirroring game. The data revealed that dyadic body competence scores positively correlate with movement synchrony, but not complexity, and that dyadic body perception scores are not associated with movement synchrony or complexity. Movement synchrony was greater when the more experienced member of the dyad was responsible for copying movements. Finally, movement synchrony and complexity were stable across the duration of the mirror game. These findings show that movement synchrony is sensitive to the composition of the dyad involved, specifically the dyad&#x2019;s embodiment, illuminating the value of dyadic approaches to understanding body movements in social contexts.</p>
</abstract>
<kwd-group>
<kwd>embodiment</kwd>
<kwd>motor synchrony</kwd>
<kwd>mirror game</kwd>
<kwd>body competence</kwd>
<kwd>body perception</kwd>
<kwd>kinematics</kwd>
<kwd>perception-action coupling</kwd>
</kwd-group>
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<fig-count count="4"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="74"/>
<page-count count="10"/>
<word-count count="7750"/>
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<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Cognitive Neuroscience</meta-value>
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</front>
<body>
<sec sec-type="intro" id="sec1">
<label>1</label>
<title>Introduction</title>
<p>As social beings, humans synchronize their behaviors with those exhibited by other humans (<xref ref-type="bibr" rid="ref38">Lakin et al., 2003</xref>; <xref ref-type="bibr" rid="ref59">Rennung and G&#x00F6;ritz, 2016</xref>; <xref ref-type="bibr" rid="ref70">Vicaria and Dickens, 2016</xref>). Humans match each other&#x2019;s limb and trunk movements, facial expressions, emotional reactions, and even vocal patterns (<xref ref-type="bibr" rid="ref7">Borrie and Delfino, 2017</xref>; <xref ref-type="bibr" rid="ref36">Lahnakoski et al., 2020</xref>; <xref ref-type="bibr" rid="ref72">Yokozuka et al., 2021</xref>; <xref ref-type="bibr" rid="ref63">Shamay-Tsoory, 2022</xref>). Inside bodies, involuntary synchrony can emerge between the hearts, endocrine systems, and brains of the self and other (<xref ref-type="bibr" rid="ref21">Feldman, 2017</xref>; <xref ref-type="bibr" rid="ref50">Nummenmaa et al., 2018</xref>; <xref ref-type="bibr" rid="ref53">Parkinson et al., 2018</xref>; <xref ref-type="bibr" rid="ref41">Mayo and Gordon, 2020</xref>; <xref ref-type="bibr" rid="ref20">De Hamilton, 2021</xref>). Generally speaking, synchrony between people is believed to support social learning and social alignment (<xref ref-type="bibr" rid="ref3">Bandura and Adams, 1977</xref>; <xref ref-type="bibr" rid="ref23">Fuhrmann et al., 2014</xref>; <xref ref-type="bibr" rid="ref64">Shamay-Tsoory et al., 2019</xref>; <xref ref-type="bibr" rid="ref18">Crone et al., 2021</xref>; <xref ref-type="bibr" rid="ref52">Pan et al., 2021</xref>) which can, in turn, strengthen social bonds (<xref ref-type="bibr" rid="ref38">Lakin et al., 2003</xref>). The social benefits of <italic>movement synchrony</italic>, i.e., matching other people&#x2019;s body movements, have been investigated in depth, and the rather colloquial consensus is that movement synchrony functions as a form of &#x2018;social glue&#x2019; (<xref ref-type="bibr" rid="ref38">Lakin et al., 2003</xref>). This consensus is built on a wealth of studies demonstrating that movement synchrony fosters feelings of social closeness or togetherness, positive affect, and prosociality (<xref ref-type="bibr" rid="ref14">Chartrand and Lakin, 2013</xref>; <xref ref-type="bibr" rid="ref59">Rennung and G&#x00F6;ritz, 2016</xref>; <xref ref-type="bibr" rid="ref70">Vicaria and Dickens, 2016</xref>; <xref ref-type="bibr" rid="ref47">Mogan et al., 2017</xref>; <xref ref-type="bibr" rid="ref73">Zampella et al., 2020</xref>). Additional evidence suggests that inducing movement synchrony through synchronizing activities, such as the mirror game, can be operationalized to enhance cognitive performance (<xref ref-type="bibr" rid="ref33">Keisari et al., 2022</xref>; <xref ref-type="bibr" rid="ref45">Moffat et al., 2024</xref>).</p>
<p>The level of movement synchrony between two people playing the mirror game varies substantially across pairs of people (dyads, henceforth; <xref ref-type="bibr" rid="ref49">Noy et al., 2011</xref>; <xref ref-type="bibr" rid="ref58">Ravreby et al., 2022</xref>; <xref ref-type="bibr" rid="ref45">Moffat et al., 2024</xref>). From a logical standpoint, this variability is to be expected, given that dyads are composed of unique individuals with their own personality traits, social experiences and bodily competencies. In terms of the consequences of such individual differences for social interactions, dyads who achieve greater movement synchrony during the mirror game are more likely to experience positive affect (<xref ref-type="bibr" rid="ref68">Tschacher et al., 2014</xref>), &#x2018;like&#x2019; each other (<xref ref-type="bibr" rid="ref58">Ravreby et al., 2022</xref>) and enjoy the interaction more (<xref ref-type="bibr" rid="ref36">Lahnakoski et al., 2020</xref>) compared to dyads whose movements are less synchronous. Moreover, the probability of dyads liking each other has been demonstrated to be closely linked to the complexity and novelty of the movements produced during the mirror game: Greater synchrony and liking of one&#x2019;s partner ensued from mirror-game movements that were more complex and novel (<xref ref-type="bibr" rid="ref58">Ravreby et al., 2022</xref>). <xref ref-type="bibr" rid="ref58">Ravreby et al.&#x2019;s (2022)</xref> work provides initial insights into the kinematic features underpinning the development of social bonds and takes important steps in considering liking from a mutual, <italic>dyadic</italic>, perspective (i.e., using ratings of liking averaged across dyads). This work represents a positive start to exploring dyadic perspectives for examining the emergence and maintenance of movement synchrony. Dyadic measures, which have been overlooked to date, give context about the makeup of the dyad <italic>per se</italic>. For example, dyadic measures of personality traits, and measures of bodily experiences/expertise that capture dyad-level differences in embodiment may be particularly informative for understanding the emergence and maintenance of movement synchrony.</p>
<p>With respect to personality traits, numerous studies examine how dyadic measures of personality traits relate to social behaviors, such as prosociality and relationship quality (e.g., <xref ref-type="bibr" rid="ref28">Heerey, 2015</xref>; <xref ref-type="bibr" rid="ref5">Bolis et al., 2021</xref>). In contrast, only a handful of studies examine the link between dyadic measures of personality traits and body movements during social interactions between two humans (<xref ref-type="bibr" rid="ref12">Carlson et al., 2019</xref>; <xref ref-type="bibr" rid="ref36">Lahnakoski et al., 2020</xref>; <xref ref-type="bibr" rid="ref2">Arellano-V&#x00E9;liz et al., 2023</xref>) and also between humans and non-human agents (i.e., robots; <xref ref-type="bibr" rid="ref1">Aly and Tapus, 2013</xref>). <xref ref-type="bibr" rid="ref2">Arellano-V&#x00E9;liz et al. (2023)</xref> recently demonstrated that dyadic measures of openness and agreeability may predict the degree of movement synchrony during conversation. Specifically, the authors found that greater agreeableness and extraversion within dyads contributed to increased movement synchrony, and greater agreeableness also contributed to increased movement complexity. <xref ref-type="bibr" rid="ref12">Carlson et al. (2019)</xref> measured dancing dyads&#x2019; trait empathy and found that dyadic levels of empathy were not related to phase locking of dance movements. The authors did, however, demonstrate that observers perceived dance movements performed by dyads composed of two individuals with high empathy scores as less similar than movements performed by dyads comprising one or more individuals with a low empathy score (<xref ref-type="bibr" rid="ref12">Carlson et al., 2019</xref>). These findings demonstrate the relevance of dyad-level differences for gaining insight into the emergence of movement synchrony. Given that very few studies, to our knowledge, have taken this approach, further exploration into the makeup of dyads is necessary to continue developing our understanding of how and why movement synchronizing activities, such as those performed during the mirror game, offer social benefits.</p>
<p>Beyond personality traits and ratings of &#x2018;liking&#x2019; one&#x2019;s synchronized partner, we propose that individual differences in embodiment are likely to help explain the emergence of movement synchrony, and perhaps movement complexity, in synchronizing activities like the mirror game. We base this proposal on research showing that expert movement improvisors and trained dancers, i.e., individuals with very high levels of embodiment acquired through intensive and prolonged training, are able to perceive subtle details in movements which allow them to synchronize with others with ease (<xref ref-type="bibr" rid="ref49">Noy et al., 2011</xref>; <xref ref-type="bibr" rid="ref30">Honisch, 2012</xref>; <xref ref-type="bibr" rid="ref27">Hart et al., 2014</xref>; <xref ref-type="bibr" rid="ref34">Kirsch et al., 2016</xref>). Congruent evidence from brain imaging studies suggests that greater embodiment (expertise) results in greater activation of a network of brain regions believed to support the simulation of observed actions (<xref ref-type="bibr" rid="ref19">Cross et al., 2006</xref>; <xref ref-type="bibr" rid="ref32">Jola et al., 2012</xref>; <xref ref-type="bibr" rid="ref34">Kirsch et al., 2016</xref>; <xref ref-type="bibr" rid="ref61">Rizzolatti and Sinigaglia, 2016</xref>). Moreover, embodiment shapes the way members of the general population engage in social interactions, potentially involving movement synchrony (<xref ref-type="bibr" rid="ref25">Garrod and Pickering, 2009</xref>; <xref ref-type="bibr" rid="ref54">Quesque and Coello, 2015</xref>) and appreciating synchrony when observing dyadic movements (<xref ref-type="bibr" rid="ref46">Moffat and Cross, 2024</xref>). Considering the evidence from expert movers and the general public, it stands to reason that the level of bodily experience or expertise possessed by members of a dyad might shape their ability to achieve and maintain synchrony, as well as the complexity of their movements while doing so.</p>
<p>In the present study, we focus on the extent to which dyadic levels of embodiment predict movement synchrony and complexity in the mirror game. More specifically, we explore how dyadic measures of beliefs regarding one&#x2019;s ability to complete physical activities (e.g., body competence; <xref ref-type="bibr" rid="ref44">Miller et al., 1981</xref>) and sensitivity to internal bodily signals (e.g., body perception; <xref ref-type="bibr" rid="ref10">Cabrera et al., 2018</xref>) are related to movement synchrony and complexity during the mirror game. Measures of body competence and body perception capture embodiment from opposing perspectives. Body competence scores offer a subjective index of one&#x2019;s cumulative experience engaging in physical activity, which is likely to correlate positively with levels of movement synchrony and complexity. Greater body perception scores are suggested to reflect an inward form of embodiment, which may detract from perception of subtle body movements performed by others and social interactions more generally (<xref ref-type="bibr" rid="ref71">Wiebking et al., 2010</xref>; <xref ref-type="bibr" rid="ref40">Lumsden et al., 2012</xref>; <xref ref-type="bibr" rid="ref16">Ciaunica et al., 2023</xref>; <xref ref-type="bibr" rid="ref17">Critchley et al., 2023</xref>; <xref ref-type="bibr" rid="ref46">Moffat and Cross, 2024</xref>). That is, greater attention to internal bodily signals may be associated with reduced levels of synchrony (<xref ref-type="bibr" rid="ref40">Lumsden et al., 2012</xref>) and movement complexity while participating in a synchronizing activity (<xref ref-type="bibr" rid="ref2">Arellano-V&#x00E9;liz et al., 2023</xref>). A deeper understanding of the role of embodiment in achieving and maintaining movement synchrony holds implications for understanding the emergence and maintenance of spontaneous movement synchrony in collaborative daily activities (i.e., passing items over a barrier), performing arts (i.e., learning dance choreography), leisure activities (i.e., in a yoga class, or during contact improvisation), and potentially, interactions with non-human agents (i.e., social robots for companionship).</p>
<p>We set out to explore the extent to which dyadic levels of embodiment (i.e., body perception and competence scores averaged within dyads) are associated with the synchrony and complexity of dyads&#x2019; movements across the course of a mirror game. We preregistered the following exploratory hypotheses on the Open Science Framework:<xref ref-type="fn" rid="fn0001"><sup>1</sup></xref> Movement synchrony will be positively correlated with dyadic body competence scores and negatively correlated with dyadic body perception scores. Movement complexity will be negatively correlated with dyadic body competence scores and positively correlated with dyadic body perception scores. Further, movement synchrony and complexity may change linearly over the course of the mirror game, and this trajectory may interact with dyadic measures of body competence and/or body perception.</p>
</sec>
<sec sec-type="methods" id="sec2">
<label>2</label>
<title>Methods</title>
<sec id="sec3">
<label>2.1</label>
<title>Participants</title>
<p>We used data from 28 participants (12 female, 16 male, mean age 21.14&#x2009;&#x00B1;&#x2009;5.08&#x2009;years) that were collected for a previous study, where participants played the mirror game with an experimental confederate before completing a cognitive task paired with brain imaging (<xref ref-type="bibr" rid="ref45">Moffat et al., 2024</xref>). Participants were recruited from Macquarie University in Sydney, Australia. All participants met the self-reported inclusion criteria of being right-handed, aged 18&#x2013;40, having no history of head injury, neurological or psychiatric diagnoses, and not currently taking a psychopharmaceutical medication (SSRIs or Ritalin). Additionally, as the data were collected in the framework of a larger neuroimaging study, participants were included if they reported no alcohol consumption within the 12&#x2009;h prior to the study or tetrahydrocannabinol (THC) use/exposure within the 24&#x2009;h prior to the study, and not playing video games frequently (e.g., more than once a week). These were the inclusion criteria for the neuroimaging study from which these secondary data originate, wherein cognitive performance and brain activity during a response inhibition task were measured (<xref ref-type="bibr" rid="ref45">Moffat et al., 2024</xref>). In this previous study, we applied the same criteria as an existing study with a similar design (<xref ref-type="bibr" rid="ref35">K&#x00F6;nig et al., 2021</xref>).</p>
<p>Participants were matched with one of two confederates; A: female, age 21; B: female, age 30; see <xref ref-type="bibr" rid="ref45">Moffat et al. (2024)</xref> for description of counterbalancing. The confederate was introduced as a new student volunteer visiting the lab for the first time (i.e., a peer to the participant, who was equally as unfamiliar with the experiment and the experimenter). Of the 28 participants, 14 were matched with confederate A (6 female, 8 male, mean age 20.93&#x2009;&#x00B1;&#x2009;5.98&#x2009;years) and the other 14 with confederate B (6 female, 8 male, mean age 21.36&#x2009;&#x00B1;&#x2009;4.2&#x2009;years).</p>
<p>Ethical approval for the original study by <xref ref-type="bibr" rid="ref45">Moffat et al. (2024)</xref> was obtained from the Macquarie University Human Research Ethics Committee (Ref: 520221102239451). Written consent was obtained from participants before the start of the experiment. Each participant received either course credit or cash honorarium (AUD $30) for their participation.</p>
</sec>
<sec id="sec4">
<label>2.2</label>
<title>Measures of embodiment</title>
<p>Participants completed two self-report questionnaires, which we employ here as proxies for embodiment. The first, the Body Competence Questionnaire (BCQ; <xref ref-type="bibr" rid="ref44">Miller et al., 1981</xref>) assesses beliefs about one&#x2019;s body&#x2019;s ability to complete physical activities. The BCQ employs a 5-point Likert scale (0&#x2009;=&#x2009;<italic>extremely uncharacteristic</italic> and 4&#x2009;=&#x2009;<italic>extremely characteristic</italic>) with statements like &#x2018;For my size, I&#x2019;m pretty strong&#x2019; or &#x2018;I&#x2019;m capable of moving quickly&#x2019;. The second, the Body Perception Questionnaire Short Form (BPQ SF; <xref ref-type="bibr" rid="ref10">Cabrera et al., 2018</xref>), assesses the degree to which an individual attends to their internal bodily signals. This scale also employs a 5-point Likert scale (1&#x2009;=&#x2009;<italic>never</italic> and 5&#x2009;=&#x2009;<italic>always</italic>) with statements such as &#x2018;I notice myself swallowing frequently&#x2019;, &#x2018;I notice my mouth being dry&#x2019;. We then averaged each of these measures separately per dyad, to obtain a joint measure per dyad, and subsequently z-scored the values at the group level.</p>
</sec>
<sec id="sec5">
<label>2.3</label>
<title>Mirror-game procedure</title>
<p>All dyads played the mirror game for a total of 5&#x2009;min. To do so, the dyad (participant and confederate) sat facing one another (<xref ref-type="fig" rid="fig1">Figure 1</xref>). Each member took a 2.5-min turn leading the game, i.e., moving their arms spontaneously, while the other member matched their arm movements as closely as possible in space and time (<xref ref-type="bibr" rid="ref22">Feniger-Schaal et al., 2021</xref>; <xref ref-type="bibr" rid="ref58">Ravreby et al., 2022</xref>). The participant always led the first 2.5&#x2009;min, followed by the confederate leading the second 2.5&#x2009;min. This was done to allow individual participants to move spontaneously without any bias from having observed an example performed by the confederate. Dyads&#x2019; arm movements were video-recorded using two GoPro video cameras positioned on a tripod between the participants. The cameras were positioned slightly above knee height as not to obstruct the dyad&#x2019;s eye contact (<xref ref-type="fig" rid="fig1">Figure 1</xref>).</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Top: Participant (red) plays the mirror game with an experimental confederate (blue). Upper-body movements were video-recorded using a pair of GoPros, one facing each member of the dyad. Bottom: Movement synchrony and complexity were calculated using the coordinates of each person&#x2019;s joints per frame, as tracked using OpenPose software.</p>
</caption>
<graphic xlink:href="fnhum-18-1401494-g001.tif"/>
</fig>
</sec>
<sec id="sec6">
<label>2.4</label>
<title>Calculation of movement synchrony and complexity</title>
<p>We divided the 2.5-min videos (<italic>N</italic>&#x2009;=&#x2009;56, 2 per dyad, i.e., participant leading and participant following) into 10 15-s segments.</p>
<sec id="sec7">
<label>2.4.1</label>
<title>Movement synchrony</title>
<p>As described in <xref ref-type="bibr" rid="ref45">Moffat et al. (2024)</xref>, the dyad&#x2019;s mean pose similarity was obtained using estimated x and y coordinates per joint obtained from OpenPose software (<xref ref-type="bibr" rid="ref11">Cao et al., 2021</xref>). We used <xref ref-type="bibr" rid="ref45">Moffat et al.&#x2019;s (2024)</xref> R code, which was previously adapted from <xref ref-type="bibr" rid="ref9">Broadwell and Tangherlini (2021)</xref>, to estimate the Euclidean distance between all pairs of body parts for each person in a frame [i.e., calculating the distances between each combination of body parts (<italic>n</italic>&#x2009;=&#x2009;42) for the participant]. These distances were stored in a separate &#x2018;pose matrix&#x2019; per person and these pose matrices were then compared (via Laplacian procedure) for each frame to establish similarity per frame. The comparison returned estimates of movement synchrony between 0&#x2009;=&#x2009;<italic>no similarity</italic> and 1&#x2009;=&#x2009;<italic>identical</italic>. We multiplied these by 100, for comparability to <xref ref-type="bibr" rid="ref45">Moffat et al. (2024)</xref>, and calculated the mean per dyad across the 15-s segment of the video. Finally, we converted the values to z-scores to homogenize the scale with that of the embodiment measures.</p>
</sec>
<sec id="sec8">
<label>2.4.2</label>
<title>Movement complexity</title>
<p>We used sample entropy (<xref ref-type="bibr" rid="ref65">Shannon and Weaver, 1949</xref>) as a proxy for movement complexity. In research investigating the kinematics of human movement, the term &#x2018;complexity&#x2019; is used interchangeably with &#x2018;predictability&#x2019; and &#x2018;entropy&#x2019; in the literature, to describe sample entropy (<xref ref-type="bibr" rid="ref51">Orlandi et al., 2020</xref>; <xref ref-type="bibr" rid="ref58">Ravreby et al., 2022</xref>; <xref ref-type="bibr" rid="ref46">Moffat and Cross, 2024</xref>). We computed the sample entropy of the x- and y-coordinates of left and right wrists of both members of a dyad across each 15-s segment of each video using the R package pracma (version 2.4.2; <xref ref-type="bibr" rid="ref6">Borchers, 2022</xref>). Next, we averaged these values per 15-s segment. Entropy values closer to zero indicate lower complexity of the movements, whereas higher entropy values reflect increased complexity. Henceforth, we refer only to movement complexity.</p>
</sec>
</sec>
<sec id="sec9">
<label>2.5</label>
<title>Data analysis</title>
<p>To explore the influence of embodiment at the level of the dyad on movement synchrony and complexity in this secondary dataset, we used the lme4 package (version 1.1&#x2013;34; <xref ref-type="bibr" rid="ref4">Bates et al., 2015</xref>) in R (version 4.3.1; <xref ref-type="bibr" rid="ref62">RStudio Team, 2023</xref>) within the RStudio IDE (version 2023.06.1; <xref ref-type="bibr" rid="ref55">R Core Team, 2023</xref>). As per our preregistration, we built two linear mixed effects models, one focusing on movement synchrony, the other on movement complexity. We added parameters to the model incrementally, beginning with varying intercepts per participant and for the index of session per confederate, to account for potential incremental increases in expertise accrued by the confederate from one experimental session to the next. Subsequently, we added the simple parameters of dyadic body competence and dyadic body perception scores, followed by time (i.e., the index of the 15-s window in the 2.5-min recording of the mirror game). Next, we added separate 2-way interactions between time and each dyadic body competence and dyadic body perception separately.</p>
<p>Parameters were retained if they contributed substantially to explaining the variance in the data (<xref ref-type="bibr" rid="ref4">Bates et al., 2015</xref>). We preregistered that we would include the index of confederates&#x2019; sessions as a varying intercept to account for confederates&#x2019; incremental acquisition of expertise across sessions. Ultimately this variable increased the complexity of our models to a degree that rendered the parameter estimates unreliable (i.e., singular or 0 convergence warning from the lme4 R package). Thus, we excluded the index of confederates&#x2019; sessions from the models and took a simpler approach to account for the confederates&#x2019; overall expertise: We included the role of the participant as &#x2018;leader&#x2019; or &#x2018;follower&#x2019; in the mirror game. If the participant was the leader, this meant the confederate was the follower, and we could thus partial out and interpret how the confederates&#x2019; expertise might influence movement synchrony. The final model predicting movement synchrony was <italic>MovementSimilarity&#x2009;~&#x2009;1&#x2009;+&#x2009;BodyCompetence&#x2009;+&#x2009;BodyPerception&#x2009;+&#x2009;Role&#x2009;+&#x2009;(1|ID)</italic>. The final model predicting movement complexity was <italic>MovementComplexity&#x2009;~&#x2009;1&#x2009;+&#x2009;BodyCompetence&#x2009;+&#x2009;BodyPerception&#x2009;+&#x2009;Role&#x2009;+&#x2009;(1|ID)</italic>. Time was not included as a parameter in our final models, as it did not explain substantial variance in the data.</p>
<p>A note regarding statistical power &#x2013; these analyses and the associated hypotheses were explicitly preregistered as &#x2018;exploratory&#x2019;. We present our findings as starting points for hypothesis generation, to be examined in future studies with greater statistical power.</p>
</sec>
</sec>
<sec sec-type="results" id="sec10">
<label>3</label>
<title>Results</title>
<p>During the mirror game, mirroring resulted in a high degree of movement synchrony (mean&#x2009;=&#x2009;0.80, SD&#x2009;=&#x2009;0.09; possible range&#x2009;=&#x2009;0&#x2013;1). The movement complexity ranged from 0.01 to 0.30 (mean&#x2009;=&#x2009;0.07, SD&#x2009;=&#x2009;0.04; closer to zero signifies lower complexity). Dyadic body competence scores showed a relatively normal distribution, as both confederates had similar individual body competence scores (mean&#x2009;=&#x2009;8.34, SD&#x2009;=&#x2009;1.54; possible range&#x2009;=&#x2009;0&#x2013;16; top left panel in <xref ref-type="fig" rid="fig2">Figure 2</xref>). Dyadic body perception scores showed a weak bimodal distribution, resulting from the two confederates having substantially different individual body perceptions scores (mean&#x2009;=&#x2009;66.55, SD&#x2009;=&#x2009;18.64; possible range&#x2009;=&#x2009;26&#x2013;130; top right panel in <xref ref-type="fig" rid="fig2">Figure 2</xref>).</p>
<fig position="float" id="fig2">
<label>Figure 2</label>
<caption>
<p>Top row: Distribution of dyadic body competence and body perception scores. Dashed line indicates the mean. Middle row: Relationship between movement synchrony and dyadic measures of body competence and body perception. Bottom row: Relationship between movement complexity and dyadic measures of body competence and body perception.</p>
</caption>
<graphic xlink:href="fnhum-18-1401494-g002.tif"/>
</fig>
<p>Next, we set out to explore the relationships between movement synchrony and complexity during the mirror game and two measures of dyadic embodiment, i.e., body competence and body perception. We hypothesized that movement synchrony would be positively associated with dyadic body competence and negatively associated with dyadic body perception. The data confirmed the expected positive association with body competence [<italic>&#x00DF;</italic>&#x2009;=&#x2009;0.02, SE&#x2009;=&#x2009;0.01, 95% CI (0.01, 0.04), <italic>p</italic>&#x2009;&#x003C;&#x2009;0.001], but showed no evidence for the expected negative association with body perception [<italic>&#x00DF;</italic>&#x2009;=&#x2009;0.01, SE&#x2009;=&#x2009;0.01, 95% CI (&#x2212;0.002, 0.02), <italic>p</italic>&#x2009;=&#x2009;0.081; middle row of <xref ref-type="fig" rid="fig2">Figure 2</xref>].</p>
<p>Next, we anticipated that movement complexity would be positively associated with body competence and negatively associated with body perception. We found no evidence to support either of these hypothesized relationships [body competence: <italic>&#x00DF;</italic>&#x2009;=&#x2009;0.00, SE&#x2009;=&#x2009;0.01, 95% CI (&#x2212;0.10, 0.10), <italic>p</italic>&#x2009;=&#x2009;0.901; body perception: <italic>&#x00DF;</italic>&#x2009;=&#x2009;0.00, SE&#x2009;=&#x2009;0.01, 95% CI (0.10, 0.10), <italic>p</italic>&#x2009;=&#x2009;0.996; bottom row of <xref ref-type="fig" rid="fig2">Figure 2</xref>].</p>
<p>Our final hypothesis was that movement synchrony and complexity may change over the course of a prolonged mirror game, and that dyadic body competence and/or perception may interact with this change. We found no evidence for any of these simple or interactional relationships, as adding <italic>Time</italic> as a parameter in our models did not improve the explanation of the variance in the data substantially (<xref ref-type="fig" rid="fig3">Figure 3</xref>).</p>
<p>In addition to the preregistered analyses above, we also explored the extent to which the mirror-game expertise of the mirror-game follower, i.e., the person copying their partner&#x2019;s movements, may influence the overall level of movement synchrony or complexity (<xref ref-type="fig" rid="fig4">Figure 4</xref>). We compared levels of movement synchrony and complexity when the mirror game was led by the participant (for whom the mirror game is novel) vs. the confederate (who has played the mirror game multiple times). We found that movement synchrony was greater when the confederate matched the participants&#x2019; movements as opposed to when the participant matched the confederates&#x2019; movements [participant following - participant leading: <italic>&#x00DF;</italic>&#x2009;=&#x2009;&#x2212;0.04, SE&#x2009;=&#x2009;0.01, 95% CI (&#x2212;0.05, &#x2212;0.03), <italic>p</italic>&#x2009;&#x003C;&#x2009;0.001]. That is, movement synchrony was greater when the expert followed the non-expert&#x2019;s movements. Further exploratory analyses revealed expertise accrued by confederates across sessions contributed significantly to explaining variance, when treated as a simple fixed effect in combination with leading or following role. Confederate session was positively associated with level of synchrony [<italic>&#x00DF;</italic>&#x2009;=&#x2009;0.01, SE&#x2009;=&#x2009;0.01, CI (0.00, 0.01), <italic>p</italic>&#x2009;=&#x2009;0.031; middle pannel of <xref ref-type="fig" rid="fig4">Figure 4</xref>]. For movement complexity, the participant&#x2019;s role did not explain substantial variance in the data and did not meet our criteria for inclusion in the model. As a result, we have no evidence for a relationship between movement complexity and the participant&#x2019;s role in the mirror game.</p>
<fig position="float" id="fig3">
<label>Figure 3</label>
<caption>
<p>Development of movement synchrony and over the course of the 2.5-min mirror game, in discrete 15-s windows.</p>
</caption>
<graphic xlink:href="fnhum-18-1401494-g003.tif"/>
</fig>
<fig position="float" id="fig4">
<label>Figure 4</label>
<caption>
<p>Left: Movement synchrony when the participant takes the role of the mirror-game leader and follower. Middle: Relationship between level of synchrony and confederate session (a proxy for increasing expertise) with participant as leader (lighter hue) and follower (darker hue). Right: Movement complexity when the participant takes the role of the mirror-game leader and follower.</p>
</caption>
<graphic xlink:href="fnhum-18-1401494-g004.tif"/>
</fig>
</sec>
<sec sec-type="discussion" id="sec11">
<label>4</label>
<title>Discussion</title>
<p>In this study, we took a dyad-centered approach to understanding the relationship between embodiment and the synchrony and complexity of movements that emerge while a dyad strives to maintain movement synchrony. We found that greater shared embodiment, in terms of dyadic body competence scores, was associated with greater movement synchrony, but not movement complexity. Dyadic body perception did not contribute to explaining inter-dyad differences in movement synchrony or complexity. We also observed that both movement synchrony and complexity remained stable over the course of the mirror game and found no evidence that the time course of movement synchrony and complexity was related to dyadic measures of embodiment. We remind readers that these exploratory analyses of a secondary dataset are to be interpreted with caution. Our motivation for these analyses was to stimulate hypothesis generation, and we recommend that the findings be subjected to further empirical examination with larger sample sizes.</p>
<sec id="sec12">
<label>4.1</label>
<title>Dyadic body competence and mirroring expertise enhance movement synchrony</title>
<p>We found that dyads with higher body competence scores showed greater movement synchrony during the mirror game. This finding aligns with existing research demonstrating that expert dancers and improvisors, whose livelihoods or hobbies involve creative but controlled body movements, achieve greater movement synchrony than non-experts (<xref ref-type="bibr" rid="ref49">Noy et al., 2011</xref>; <xref ref-type="bibr" rid="ref30">Honisch, 2012</xref>; <xref ref-type="bibr" rid="ref27">Hart et al., 2014</xref>; <xref ref-type="bibr" rid="ref34">Kirsch et al., 2016</xref>). Thus, our findings are consistent with existing evidence that embodiment is important for achieving synchrony. Moreover, the positive association between dyadic body competence scores and movement synchrony highlights two important phenomena. First, embodiment gained through the general public&#x2019;s (in this case, Australian undergraduate students&#x2019;) everyday activities and hobbies also contributes to the ability to achieve movement synchrony with a stranger. Second, a high degree of movement synchrony is more likely when both members of a dyad have strong positive beliefs about their body&#x2019;s ability to complete physical tasks.</p>
<p>We also observed that movement synchrony was greater when the experimental confederate was the follower rather than the leader. We initially interpreted this to mean that the level of movement synchrony is impacted differently by a dyad member&#x2019;s bodily experience with the mirror game, depending on whether they are leading or following. We are thankful to a reviewer for raising the possibility that confederates&#x2019; knowledge of the study&#x2019;s hypotheses, the fact that the movement synchrony would be evaluated, and the social benefits of movement synchrony described in the literature, may have engendered a different form of motivation than that experienced by participants. Indeed, it appears (middle pannel of <xref ref-type="fig" rid="fig4">Figure 4</xref>), that synchrony increased alongside the number of sessions completed by each experimental confederate, suggesting that confederates&#x2019; expertise does indeed play a role in the level of synchrony. However, the overall difference in the level of synchrony when the participant was leading and following is consistent across confederate sessions (i.e., no interaction), suggesting that the differing forms of motivation experienced by participants and experimental confederates might underpin the difference in synchrony between participant and confederate-led movements.</p>
<p>Considering the influence of expertise on movement synchrony first, our findings add to existing evidence from <xref ref-type="bibr" rid="ref27">Hart et al. (2014)</xref> that dyads comprising two expert improvisors, relative to two novice improvisors, can achieve greater movement synchrony in the mirror game. That is, greater embodiment is particularly beneficial for supporting the perception&#x2013;action processing involved in matching another person&#x2019;s movements in time and space (<xref ref-type="bibr" rid="ref31">Honisch et al., 2016</xref>). With greater embodiment (i.e., body competence or experience mirroring), movement synchrony may emerge more fluidly in interactions with others. This proposition aligns with evidence that embodiment, gained through physical experience, may enable more fluent simulation of observed movements, which in turn is associated with greater activation of sensorimotor brain regions while observing familiar movements (<xref ref-type="bibr" rid="ref19">Cross et al., 2006</xref>; <xref ref-type="bibr" rid="ref32">Jola et al., 2012</xref>; <xref ref-type="bibr" rid="ref34">Kirsch et al., 2016</xref>). Second, we turn to the probable influence of differing motivations on movement synchrony in synchronizing activities, such as the mirror game. This aligns with existing findings, suggesting individuals&#x2019; motivations (i.e., to bond with others) are predictive of movement synchrony levels (<xref ref-type="bibr" rid="ref42">Miles et al., 2010</xref>; <xref ref-type="bibr" rid="ref40">Lumsden et al., 2012</xref>; <xref ref-type="bibr" rid="ref24">Fujiwara et al., 2020</xref>). Moreover, the perspective that motivations shape movement synchrony highlights a potential weakness of experimental confederates for studying movement synchrony. Considered differently, the present work provides preliminary evidence suggesting that expertise with synchrony and motivation to maintain synchrony should be considered when designing studies or interventions involving synchronizing activities.</p>
<p>Moreover, our findings have useful implications for contexts where movement synchrony is operationalized to improve social connections or cognition (e.g., <xref ref-type="bibr" rid="ref57">Ramseyer and Tschacher, 2011</xref>; <xref ref-type="bibr" rid="ref69">Tun&#x00E7;gen&#x00E7; and Cohen, 2016</xref>; <xref ref-type="bibr" rid="ref33">Keisari et al., 2022</xref>; <xref ref-type="bibr" rid="ref45">Moffat et al., 2024</xref>). Greater synchrony is likely to ensue when a trained or experienced individual takes the role of the follower. We also propose that experience can be accrued, as our experimental confederates were not &#x2018;experienced&#x2019; at playing the mirror game prior to helping with the data collection, and the experience gained through data collection was sufficient for them to improve their perception-action correspondences in such a way that led to greater synchrony.</p>
</sec>
<sec id="sec13">
<label>4.2</label>
<title>Measures of embodiment do not predict complexity of dyadic movements</title>
<p>We did not find movement complexity to be modulated by any measures of embodiment (body competence/perception) or bodily experience with the mirror game. We interpret this as evidence that successful synchronization relies more on the follower&#x2019;s perception and matching of movements than the leader&#x2019;s adjustment of movement complexity. In other words, we found no evidence that dyadic or individual levels of embodiment were associated with behaviors like making simpler or more predictable movements to help the follower, or more complex, less predictable, movements as a challenge to the follower. This observation is reinforced by our findings that movement complexity remained stable over the course of the mirror game.</p>
<p>We anticipated synchrony might increase over the course of the mirror game as participants become entrained with each other&#x2019;s movements and that this may co-occur with less complex movement that could be more easily predicted by the follower. We also evaluated an alternative prediction based on <xref ref-type="bibr" rid="ref58">Ravreby et al.&#x2019;s (2022)</xref> report that novelty and complexity of mirror game movements facilitate greater interpersonal liking: we anticipated that dyads might increase the complexity of their movements over time, thereby reducing synchrony, as they may subconsciously seek more positive social experiences as the mirror game progressed. A linear increase in novelty, but not complexity is indeed visible in <xref ref-type="bibr" rid="ref58">Ravreby et al.&#x2019;s (2022)</xref> data visualizations. While our findings regarding the stability of movement synchrony and complexity are aligned with Ravreby&#x2019;s visualization, it is possible that a non-linear approach to quantifying changes in movement synchrony or complexity over time may offer more nuanced insights. With respect to synchrony, it is possible that the overall level of synchrony resulting from the mirror game is too high with too little variance between dyads from the onset of the game. In other words, our data may show a type of ceiling effect for synchrony. Another consideration is that a larger sample size would be beneficial for assessing changes over time, as the effect size of such changes might be smaller than detectable with the current sample (see Data analysis section for note on statistical power).</p>
</sec>
<sec id="sec14">
<label>4.3</label>
<title>Dyadic perspectives for illuminating dyadic interactions</title>
<p>In this study, we explicitly considered dyadic scores related to embodiment (i.e., body competence and perception) as opposed to individual scores. While information about individuals can shed light on how individuals might behave, information about dyads offers more detailed insight into dyadic behaviors that emerge in interactive social contexts (<xref ref-type="bibr" rid="ref2">Arellano-V&#x00E9;liz et al., 2023</xref>). Our findings add to the relatively sparse landscape of studies describing how dyad-level traits, experiences and skills influence the emergence of interpersonal synchrony by showing that greater dyadic embodiment enhances the level of synchrony that a dyad can achieve. We do not wish to neglect the wealth of research examining individual traits in relation to social interactions, as opposed to dyads&#x2019; traits (e.g., <xref ref-type="bibr" rid="ref37">Lakens and Stel, 2011</xref>; <xref ref-type="bibr" rid="ref43">Miles et al., 2011</xref>; <xref ref-type="bibr" rid="ref56">Ragert et al., 2013</xref>; <xref ref-type="bibr" rid="ref68">Tschacher et al., 2014</xref>, <xref ref-type="bibr" rid="ref67">2018</xref>; <xref ref-type="bibr" rid="ref8">Brambilla et al., 2016</xref>; <xref ref-type="bibr" rid="ref15">Cheng et al., 2017</xref>; <xref ref-type="bibr" rid="ref36">Lahnakoski et al., 2020</xref>; <xref ref-type="bibr" rid="ref66">Stupacher et al., 2020</xref>; <xref ref-type="bibr" rid="ref74">Zimmermann et al., 2021</xref>; <xref ref-type="bibr" rid="ref26">Glass and Yuill, 2023</xref>). Instead, we strive to highlight the value of examining dyads&#x2019; behavior using dyadic measures. Future work could, for example, further explore additional dyadic measures that pertain to embodiment, such as experience with the arts (<xref ref-type="bibr" rid="ref13">Casale et al., 2023</xref>), or even dyadic disembodiment traits (<xref ref-type="bibr" rid="ref16">Ciaunica et al., 2023</xref>).</p>
<p>Another potential avenue for future research is related to the increasing prevalence of social interactions of non-human agents (i.e., chatbots, social robots) in everyday life. Non-human agents&#x2019; embodiment, specifically the kinematics of their interactions with humans, is central to their acceptance by users (<xref ref-type="bibr" rid="ref39">Lorenz et al., 2016</xref>; <xref ref-type="bibr" rid="ref48">Nalepka et al., 2019</xref>; <xref ref-type="bibr" rid="ref60">Rinott and Tractinsky, 2022</xref>). A deeper understanding of the emergence and maintenance of movement synchrony within human&#x2013;robot dyads may help enhance perceptions, acceptance and then ensuing interactions (<xref ref-type="bibr" rid="ref29">Henschel and Cross, 2020</xref>). Existing evidence suggests that people prefer social robots with personalities resembling their own (<xref ref-type="bibr" rid="ref1">Aly and Tapus, 2013</xref>), emphasizing how dyadic approaches are equally promising for understanding interactions with non-human agents.</p>
</sec>
</sec>
<sec sec-type="conclusions" id="sec15">
<label>5</label>
<title>Conclusion</title>
<p>We set out to explore the influence of dyadic indices of embodiment (body competence and perception) on dyads&#x2019; abilities to achieve and maintain movement synchrony, and the complexity of the movements they made during the mirror game. The data revealed that higher body competence scores and mirror-game expertise, but not body perception scores, allowed dyads to achieve greater movement synchrony. Movement complexity during the mirror game was not influenced by any measure investigated here. Moreover, the data suggest that movement synchrony and complexity remain relatively stable across mirror game rounds. We conclude by emphasizing the value of dyadic approaches for uncovering the dynamics that modulate dyadic body movements. Dyadic approaches stand to enable the design of more effective movement synchrony interventions to promote social and cognitive behaviors.</p>
</sec>
<sec sec-type="data-availability" id="sec16">
<title>Data availability statement</title>
<p>Publicly available datasets were analyzed in this study. This data can be found at: <ext-link xlink:href="https://osf.io/gd5nb/" ext-link-type="uri">https://osf.io/gd5nb/</ext-link>.</p>
</sec>
<sec sec-type="ethics-statement" id="sec17">
<title>Ethics statement</title>
<p>The studies involving humans were approved by Macquarie University Human Research Ethics Committee. The studies were conducted in accordance with the local legislation and institutional requirements. The participants provided their written informed consent to participate in this study.</p>
</sec>
<sec sec-type="author-contributions" id="sec18">
<title>Author contributions</title>
<p>RM: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Project administration, Supervision, Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing. LR: Conceptualization, Formal analysis, Investigation, Methodology, Visualization, Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing. CC: Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing. EC: Conceptualization, Funding acquisition, Project administration, Resources, Supervision, Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing.</p>
</sec>
</body>
<back>
<sec sec-type="funding-information" id="sec19">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. This project was supported by a Leverhulme grant to ESC (PLP-2018-152) and the Professorship for Social Brain Sciences at ETH Zurich. LR was supported by a fellowship of the German Academic Exchange Service (DAAD). Open access funding by ETH Zurich.</p>
</sec>
<ack>
<p>We thank the confederates, Hannah Cahill and Sabrina Diep, for their involvement in the data collection.</p>
</ack>
<sec sec-type="COI-statement" id="sec20">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="disclaimer" id="sec21">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<fn-group>
<fn id="fn0001">
<p><sup>1</sup><ext-link xlink:href="https://osf.io/6rc3g" ext-link-type="uri">https://osf.io/6rc3g</ext-link>
</p>
</fn>
</fn-group>
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